Die Gattung Calceolaria (Scrophulariaceae) in Chile [The genus Calceolaria (Scrophulariaceae) in Chile]

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Language: German with bilingual abstract in English and German

This revision is the first critical work about the genus Calceolaria in Chile since Reiche’s contribution to the 1911 Flora de Chile. All the accepted species are described and the important features, usually including habit, are illustrated.

The characteristic three-dimensional flower structure of the genus coupled with the difficulty of understanding it from herbarium material has until now been a significant contributing factor to the misidentifications, uncertainties and ignorance concerning the chilean species of Calceolaria. Due to sparse collections, variation within and between populations and species, as well as distribution patterns, have hitherto remained obscure. This also goes for concepts concerning character evolution and the generic limits within this, the largest group of extra-tropical species of Calceolaria. Extensive fieldwork and cultivation of nearly all the species from approximately 400 populations were the key to the understanding of the character complexes.

In this work the chilean component of the genus comprises 50 species, approximately a fifth of the entire genus. About 74% of these are endemic to Chile, two taxa occur also in Peru, and a further 16 spread across into Argentina.

One new species C. flavovirens is described. Ten species express clear morphologically related regional differentiation. As a result, as well as the type-subspecies, 19 further subspecies are recognised. Two of these subspecies are newly described here, namely C. densifolia subsp. laxa, and C. corymbosa subsp. santiagina.

The species were subdivided as follows:
- C. corymbosa with C. corymbosa subsp. corymbosa, C. corymbosa subsp. floccosa, C. corymbosa subsp. mimuloides, C. corymbosa subsp. montana, C. corymbosa subsp. santiagina and C. corymbosa subsp. tetraphylla
- C. meyeniana with C. meyeniana subsp. meyeniana, C. meyeniana subsp. cheiranthoides, C. meyeniana subsp. glabrata and C. meyeniana subsp. nahuelbutae
- C. ascendens with C. ascendens subsp. ascendens, C. ascendens subsp. glandulifera and C. ascendens subsp. pristiphylla
- C. collina with C. collina subsp. collina, C. collina subsp. conferta and C. collina subsp. subincisa
- C. dentata with C. dentata subsp. dentata, C. dentata subsp. araucana and C. dentata subsp. cummingiana
- C. arachnoidea with C. arachnoidea subsp. arachnoidea and C. arachnoidea subsp. nubigena
- C. filicaulis with C. filicaulis subsp. filicaulis and C. filicaulis subsp. luxurians
- C. valdiviana with C. valdiviana subsp. valdiviana and C. valdiviana subsp. crassifolia
- C. glandulosa with C. glandulosa subsp. glandulosa and C. glandulosa subsp. alicahuensis
- C. densifolia with C. densifolia subsp. densifolia and C. densifolia subsp. laxa.

A particularly difficult complex is formed by the taxa within C. integrifolia. These are not treated here, and together with C. integrifolia (in the strictest sense) are considered as C. integrifolia s.l.

The chilean species of Calceolaria colonise, with the exception of the arid areas in Northern Chile, all habitats. They are found from the coast to the Andes and from 10 m a.s.l. to over 4000 m a.s.l. in all the administrative Regions of Chile. The centre of diversity of the genus has one locus, frorm Region IV over the "Zona Central“ with Regions V, VI, VII, and VIII to the south of mediterranean Chile. To the north the species number rapidly declines, with taxa inhabiting only special areas of the coastal ranges and the high Andes. Contrarily, to the South, the species number slowly drops off, as the taxa that are characteristically distributed into Argentina play a more significant role.

Growth habit and inflorescences are greatly influenced by altitude and climate. Both characters show a continuous spectrum of morphological change. The variety in growth habit, at its greatest extremes, changes from small annual rosette plants, to rosulate or subrosulate perennials, to woody bushes and shrubs.

The inflorescences are organised as a type of open thyrse. They exhibit a complete range of forms from a luxuriant synflorescence constructed of a main florescence and numerous paraclades, to a single-flowered reduced synflorescence. The particular structure and variability of the synflorescence is presented for every species.

The reticulate surface structure of the seeds was found to be uniform across the taxa and was not useful for species groupings. The variation in the seeds bore no correlation to the system based on flower characters proposed here.

Together with new chromosome counts, the chromosome numbers of nearly 80% of all chilean species are now knownn. They show that, contrary to the tropical species, the Chilean taxa are almost exclusively diploid (2n = 18). Polyploids are found increasingly towards the south, in the Patagonian and Tierra del Fuego taxa, and also to the north in the tropical regions.

The flowers of the chilean Calceolaria species possess many varied and good characters for taxon delimitation. In most cases the species can be characterised by their specific flower forms alone. The colours of the flowers and their markings, the upper lip form, the relationship of the upper lip to the lower, the width and orientation of upper lip opening, and the form of the lower lip, depression of the upper side of the lower lip, as well as the masking, width and form of the lower lip opening provide a wealth of definitive and, above all, constant characters. The ratio of anther to filamlent length, form and length of the glandular lobe, margin of the glandular field, together with the length of the glandular hairs, and the form of the glands themselves are also of taxonomic significance.

All these differences are undoubtedly related to the pollination biology, but this requires much more investigation.

The classification of the Chilean species is essentially based on flower characters. The chilean representatives are divided into four sections, one of which, the monospecific section Tenella, is newly described. Section Kremastocheilos Witasek contains the Patagonian C. uniflora, and section Calceolaria the northern Chilean C. pinnata (the type section has its greatest species concentration in the tropical Andes). The majority, a comparatively broadly homogeneous group of species is placed in section Cheiloncos Witasek. It is subdivided into 14 informal Greges supported by characters from growth habit and inflorescence structure.

The 14 Greges are roughly divided into two large groups based on floral morphology. Group A is defined by having an upper lip one third the length of the lower lip and anthers longer than the filaments. Group B is defined by having an upper lip at least one third the length of the lower lip and anthers distinctly shorter than the filaments.

This work clearly demonstrates that a classification of the genus based on 3 subgenera (as suggested by Molau, 1988), two of which are extra-tropical and therefore also include the Chilean species, cannot be supported.

Contents

Abstract VI 

Zusammenfassung VII 

I. Einleitung 1 

II. Material und Methoden 2 

III. Die vegetativen Merkmale und ihre systematische 

Bedeutung 3 

1. Die Wuchsformen 3 

1.1. Wuchsform und Infloreszenz 4 

1.2 Merkmalsphylogenie der Wuchsformen 6 

1.3 Die verschiedenen Wuchsformen 7 

1.4 Systematische Bedeutung der Wuchsformen 17 

2. Das Indument 17 

3. Die Blätter 17 

4. Die Wurzeln 18 

IV. Die generativen Merkmale und ihre systematische 

Bedeutung 18 

1. Die Infloreszenzen 18 

1.1. Die Synfloreszenz und ihre Abgrenzung 19 

1.1.1. Typologische Charakterisierung 19 

1.1.2. Abgrenzung der Synfloreszenz 19 

1.2. Differenzierung der Parakladien 19 

1.2.1 Basitone Forderung 19 

1.2.2 Akrotone Forderung 19 

1.3. Die Partialfloreszenzen (Cymen) 22 

1.4. Merkmalsplastizität 23 

1.5. Die Merkmale und Merkmalsreihen 24 

1.5.1. Die Merkmale 24 

1.5.2. Die Merkmalsreihen 24 

1.6. Die Synfloreszenzen der einzelnen Arten 28 

1.7. Die Synfloreszenz als systematisches Merkmal 38 

2. Die Büte 38 

2.1. Der Kelch 40 

2.2. Die Krone 40 

2.2.1. Die Oberlippe 40 

2.2.2. Die Unterlippe 41 

2.2.3. Der Drüsenlappen, die DrDsenfelder (Elaiophore) und Drüsenhaare 42 

2.2.3a. Der Drüsenlappen 42 

2.2.3b. Das Drüsenfeld 43 

2.2.3c. Die Drüsenhaare 44 

2.2.3d. Korrelationen zwischen den Einzelmerkmalen 44 

2.2.4. Die Zeichnung der Blüte 45 

2.3. Die Staubblätter 47 

2.3.1. Langenverhältnis Filament/Anthere 47 

2.3.2. Thekenform, Ausrichtung und Öffnung der Theken 47 

2.3.3. Stellung der Staubblätter zum Griffel 47 

2.4. Das Gynoeceum und die Kapsel 48 

3. Der Pollen 49 

4. Der Same 49 

4.1. Morphologie 49 

4.1.1. Größe und Form 49 

4.1.2. Farbe 50 

4.1.3. Ornamentierung 50 

4.1.4. Variabilität in der Ornamentierung 50 

4.2. Die Samenmorphologie als systematisches Merkmal 51 

V. Bestaubung und Hybridisierung 54 

VI. Cytologie 54 

VII. Verbreitung und Okologie 56 

1. Die Verbreitung von Calceolaria in Chile 56 

2. Verbreitungsmuster im Bereich der Laguna del 
Maule und der Nevados de Chillan 59 

3. Standorte 61 

VIII. Die Gruppierung der chilenischen Arten der Gattung Calceolaria - 
Auswirkungen auf eine zu revidierende Gattungsgliederung 63 

1. Historischer Überblick 63 

2. Kritische Prilfung des aktuellen Gliederungsmodells 63 

2.1. Die Gattungsgliederung nach Molau 63 

2.2. Bewertung der untergliedernden Merkmale nach 
Analyse der chilenischen Arten 64 

2.2.1. Wuchsform 64 

2.2.2. Position des Fruchtknotens 64 

2.2.3. Verhältnis der Filamentlange zur Antherenlänge 64 

2.2.4. Samenoberflächenstruktur 65 

2.2.5. Ploidiestufe 65 

2.3. Resumée 65 

3. Die Gliederung der chilenischen Arten 65 

3.1. Die Blütenmorphologie als Merkmal zur Gliederung der chilenischen 
Arten 65 

3.2. Verwandtschaftslinien und Vernetzungsmuster 
der chilenischen Arten 65 

3.3. Charakterisierung und Verknilpfung der Greges 
innerhalb der Sektion Cheiloncos 66 

3.4. Synopsis der chilenischen Arten 70 

3.5. Die Gattungsgliederung im Schema 71 

4. Bemerkungen ffir eine Gesamtgliederung 71 

IX. Schlüssel zu den Arten 77 

X Die Arten 83 

Sektion Calceolaria 84 

Sektion Kremastocheilos 86 

Sektion Tenella 89 

Sektion Cheiloncos 92 

Grex I 92 

Grex II 96 

Grex III 107 

Grex IV 126 

GrexV 161 

Grex Vl 191 

Grex VII 194 

Grex VIII 197 

Grex IX 205 

Grex X 207 

Grex XI 240 

Grex XII 247 

Grex XIII 256 

Grex XIV 263 

XI. Zweifelhafte Namen 277 

XII. Dank 277 

XIII. Literatur 278 

XIV. Index der Taxa 282

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